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Research Article
Cirsium tatakaense (Compositae), a new species from Taiwan
expand article infoChih-Yi Chang‡, Hsy-Yu Tzeng‡, Yen-Hsueh Tseng‡
‡ National Chung-Hsing University, Taichung, Taiwan

Corresponding author: Yen-Hsueh Tseng ( tseng2005@nchu.edu.tw )

Academic editor: Anthony R. Magee
© 2019 Chih-Yi Chang, Hsy-Yu Tzeng, Yen-Hsueh Tseng.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Chang C-Y, Tzeng H-Y, Tseng Y-H (2019) Cirsium tatakaense (Compositae), a new species from Taiwan. PhytoKeys 117: 119-132. https://doi.org/10.3897/phytokeys.117.29380
Open Access

Abstract

A new species of Cirsium, Cirsium tatakaense Y.H.Tseng & C.Y.Chang, from central-southern Taiwan is described and illustrated. This species is similar to C. kawakamii Hayata in leaf shape, achene and chromosome number (2n = 64), but can be readily distinguished from C. kawakamii by the narrower leaf lobes, usually higher number of florets and phyllaries, the purplish-red corolla (vs. white) and larger pollen grains. A key to the species of Cirsium in Taiwan is also presented.

Keywords

New species, Chromosome number, Cirsium tatakaense , Cirsium kawakamii , Compositae , Taiwan

Introduction

Cirsium Mill. is a genus of Compositae and comprises approximately 250 species widely distributed throughout the world’s temperate and subtropical zones (Kadota 2007). A total of 46 species are native to China (Shih and Greuter 2011), 64 to Japan (Iwatsuki et al. 1995) and 9 to Taiwan (Peng 2003).

Kitamura (1937) established the first infrageneric classification for East Asian Cirsium. In this classification, the Taiwanese species were placed in two of the sections, sect. Pseudoeriolepis (Nakai) Kitam. and sect. Onotrophe (Cass.) DC. and five subsections, subsect. Arenicola Kitam., subsect. Australicirsium Kitam., subsect. Nipponocirsium Kitam., subsect. Sinocirsium Kitam. and subsect. Spanioptilon Less. The latter subsection was subsequently raised to the rank of section (sect. Spanioptilon (Less.) Shih) by Shih (1984) and maintained by Iwatsuki et al. (1995).

Recently, we discovered a rare Cirsium growing in the high mountain areas of central-southern Taiwan. Based on the systems of Kitamura (1937) and Iwatsuki et al. (1995), this taxon belonged to section Onotrophe subsection Nipponocirsium and appeared to be similar to Cirsium kawakamii Hayata, sharing the longer leaf lobes, nodding bowl-shaped mature capitula and corolla lobes as long as the inflated part of the corolla tube.

Materials and methods

We compared the new species to the other two species of Cirsium sect. Onotrophe in Taiwan.

Herbarium examination

Materials primarily comprised fresh and dried specimens; voucher specimens were deposited in TCF, PPI and TNM. The herbaria referenced included HAST, KYO, PPI, TAI, TAIF, TCF, TI and TNM.

Pollen morphology

Pollen grains, all from fresh material, were directly mounted on a stub without any pretreatment and sputter coated with gold (Quorum SC7620) for observation with a scanning electron microscope (Hitachi S-3400N). The shape, size and exine ornamentation were studied following Erdtman (1952) and Hesse et al. (2009). Vouchers for the pollen material studied are provided in Table 1.

Table 1.
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Voucher material for the Cirsium Mill. pollen morphology.

Taxa Location Coordinate Altitude Date Voucher
C. tatakaense TAIWAN. Kaohsiung City Tauyuan District, Kuaiku to Yakou 23°16.03'N, 120°56.24'E 2600 m 21 Oct 2014 C. Y. Chang, 160 (TNM)
TAIWAN. Nantou County Sinyi Township, Tataka 23°29.24'N, 120°52.71'E 2620 m 15 Nov 2014 C. Y. Chang 182 (TCF)
TAIWAN. Nantou County Sinyi Township, Tataka to Shihshan 23°28.56'N, 120°52.74'E 2530 m 26 Oct 2015 C. Y. Chang, 842 (TNM)
TAIWAN. Nantou County Sinyi Township, Tataka to Shihshan 23°29.24'N, 120°52.71'E 2631 m 22 Dec 2015 C. Y. Chang 1017 (TCF)
C. kawakamii TAIWAN. Taichung City Heping District, Mt. Pintian to Mt. Dabajian 24°26.52'N, 121°15.83'E 2800 m 13 Aug 2014 C. Y. Chang 264 (TNM)
TAIWAN. Taichung City Heping District, Mt. Syue Trail 8.9 km 24°23.53'N, 121°14.20'E 3500 m 3 Oct 2015 C. Y. Chang 774 (TCF)
TAIWAN. Nantou County Ren’ai Township. Nengao cross-ridge historic trail 11 km 24°2.79'N, 121°15.77'E 2600 m 29 Jan 2018 C. Y. Chang 1605 (TNM)
C. arisanense TAIWAN. Taichung City Heping District, Mt. Syue trail 8.9 km 24°23.60'N, 121°13.98'E 3450 m 3 Sept 2015 C. Y. Chang 756 (TCF)
TAIWAN. Nantou County Ren’ai Township, Rueiyan river pipes road 2 km 24°06.95'N, 121°11.96'E 2240 m 27 May 2016 C. Y. Chang 1275 (TCF)

Karyotype analysis

Karyotype analysis was performed by following the same procedure applied by Ozcan et al. (2011) and Yüksel et al. (2013). Root tips were collected on sunny mornings and preserved in 0.002 M 8-hydroxyquinoline solution below 10 °C for eight hours. This material was then fixed with Carnoy’s solution (1 part acetic acid: 3 parts EtOH) for at least 24 hours at 4 °C. The fixed roots were then stained with acetic-orcein for 24 hours at room temperature, squashed and the slides examined using an stereo microscope (ACCU-ScoPE 3025). Voucher material is presented in Table 2.

Table 2.
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Voucher material for the Cirsium Mill. karyotype analysis.

Taxa Location Coordinate Altitude Date Voucher
C. tatakaense TAIWAN. Chiayi County Alishan Township, Tataka to Paiyun lodge 24°28.40'N, 120°54.23'E 2800 m 15 May 2016 C. Y. Chang 1269 (TCF)
C. kawakamii TAIWAN. Taichung city Heping District, Mt. Syue trail 8.9 km 24°23.73'N, 121°13.94'E 3371 m 8 Nov 2015 C. Y. Chang 874 (TCF)
TAIWAN. Taichung city Heping District, Mt. Syue trail 8.9 km 24°23.73'N, 121°13.94'E 3371 m 21 May 2016 C. Y. Chang 1271 (TCF)

Results and discussion

Morphological comparison

Following Kitamura (1937) and Iwatsuki et al. (1995), Cirsium tatakaense is placed in section Onotrophe together with C. kawakamii and C. arisanense. Amongst them, C. arisanense belongs to subsection Australicirsium, which is characterised by having rosette leaves, pot-shaped capitula which are erect or nodding when mature, corolla lobes equal in length to the inflated part of corolla tube and corona-like achene beaks. Both C. tatakaense and C. kawakamii belong to subsection Nipponocirsium, which is characterized by not having rosette leaves, larger bowl-shaped capitula which are nodding when mature, corolla lobes equal in length to the inflated corolla tube and tube-like achene beaks. In comparison with C. kawakamii (Table 3), C. tatakaense has purplish corolla (vs. white in C. kawakamii) (Figure 5[2]), usually more florets (136)161−308 (vs. (61)115−222 in C. kawakamii) (Figure 5[2]) and phyllaries 111−199 (vs. 79−123 in C. kawakamii) (Figure 5[2]) and narrower leaf lobes 7.3−11.9 mm (vs. 17.2−18.6 mm in C. kawakamii) (Figure 5[1]).

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Summary of characters between the species of Cirsium sect. Onotrophe in Taiwan.

Characters C. tatakaense C. kawakamii C. arisanense
Leaf size (cm) 27.2−34.8 cm × 16.4−19.4 cm 27.5−30.6 cm × 17.8−20.2 cm 10.6−21.3 cm × 3.0−6.3 cm
Leaf shape Elliptic to broadly elliptic Elliptic to broadly elliptic Narrowly elliptic to deltoid
Leaf margin Mainly pinnatisect Pinnatisect or bipinnatisect Pinnatipartite or bipinnatipartite
Leaf lobes Size 6.4−7.5 cm × 7.3−11.9 mm 8.3−10.4 cm × 17.2−18.6 mm 0.7−2.9 cm × 6.6−16.0 mm
Pair of leaflobes 4−6 6−7 6−10
Mature Capitula Nodding Nodding Erect or nodding
Involucre shape Bowl-shaped (upper width ≥ base) Bowl-shaped (upper width ≥ base) Pot-shaped (upper width< base)
Corolla colour Purplish-red White Yellow
Floret number (136)161−308 (61)115−222 87−133
Phyllary number 111−199 79−123 93−114
Beak of achene Tube-like Tube-like Corona-like
Pollen size (P/E) 34.2−42.6μm/ 35.2−44.7 μm 31.7−34.5 μm/ 34.3−37.1 μm 41.7−51.0 μm/ 44.7−49.3 μm
Pollen spine base width 4.2−5.6 μm 2.0−2.3 μm 2.8−4.8 μm
Chromosome number 2n = 64 2n = 64 2n = 34 (Peng and Hsu 1978)
Distribution Endemic to Taiwan; open areas of fog forests at 2000−3000 m alt. central-southern Taiwan (Fig. 3) Endemic to Taiwan; gullies and valleys at 1500−3500 m alt. central-northern Taiwan (Fig. 3) Endemic to Taiwan; widely distributed in open areas of mountain area at 1500−3800 m alt. (Fig. 3)

Chromosome number

The basic number of chromosomes amongst Cirsium species is often 2n = 34 (Hsu 1970; Funk et al. 2009; Chen and Yeh 2010a; Chen and Yeh 2010b), including in C. arisanense (Peng and Hsu 1978). However, the chromosome number of C. tatakaense is 2n = 64 (Fig. 6A), which is the same as that of C. kawakamii (Fig. 6B), indicating that the two species are similar in this respect. Notably, other taxa of the same subsection in Japan are 2n = 68 (Iwatsuki et al. 1995). These findings imply that subsect. Nipponocirsium are tetraploids with aneuploid cells.

Palynological study

Cirsium tatakaense pollen has a larger diameter, up to 36−43 μm (vs. 32−35 μm in C. kawakamii) and its surface spines have broader bases of 4.2−5.6 μm (vs. 2.0−2.3 μm in C. kawakamii). The pollen grains of C. tatakaense are similar to C. arisanense (Fig. 7A, C). However, C. kawakamii (similar to C. tatakaense in macroscopic morphology) has the smallest pollen grains and spine in Taiwan (Fig. 7B). Pollen morphology is associated with pollination, thus implying reproductive isolation between the two species.

Comparison of the distribution between C. tatakaense and C. kawakamii

Compared with C. tatakaense, C. kawakamii occurs at higher altitudes (up to 3500 m); C. tatakaense is seldom discovered over altitudes of 3000 m. In addition, C. kawakamii is usually distributed in alpine gullies and valleys, whereas C. tatakaense often appears on spacious roadsides, seemingly with no preference for valley habitats. Therefore, we believe that C. kawakamii prefers shaded and moist environments, whereas C. tatakaense prefers open areas with higher drought tolerance. Some geographical segregation appears to exist in the distributions of C. tatakaense and C. kawakamii.

Taxonomic treatment

Key to the species of Cirsium Mill. in Taiwan

1 Biennial herb; involucre tube-shaped (length 2 times than width); corolla lobes < 2.5 mm long C. ferum
– Perennial herb; involucre pot or bowl-shaped (length approximates width), corolla lobes > 2.5 mm long 2
2 All leaves cauline, basal rosette leaves absent 3
– Leaves in both a basal rosette as well as cauline 5
3 Leaves densely cobwebbed on abaxial surface; mature capitula erect, involucre pot-shaped (upper width shorter than base); apical parts of inner phyllaries inflated, obtuse; outer phyllaries lanceolate, apex acute without spine; corolla lobes obviously longer than the inflated part of corolla tube C. lineare
– Leaves glabrous on both surfaces; mature capitula nodding, involucre bowl-shaped (upper width greater or equal to base); apical parts of inner phyllaries acute or acuminate; outer phyllaries elliptic with long spine at the apex; corolla lobes as long as the inflated part of corolla tube 4
4 Corollas white; leaves pinnatisect or bipinnatisect, lobes > 15 mm wide C. kawakamii
– Corollas purple; leaves mainly pinnatisect, lobes < 12 mm wide C. tatakaense
5 Phyllaries narrowly ovate 6
– Phyllaries subulate 7
6 Corollas white; phyllaries lanceolate, inner and outer phyllaries similar in length; stems cauline, without rhizome C. brevicaule
– Corollas purple; phyllaries narrowly ovate to ovate, inner and outer phyllaries distinct in length; stems both cauline and rhizomatous C. morii
7 Apical prominently parts of phyllaries longer than 4 mm, blade-like; corolla lobes as long as the inflated part of corolla tube 8
– Apical prominently parts of phyllaries shorter than 4 mm, spine-like; corolla lobes shorter than the inflated part of corolla tube 9
8 Leaf abaxial surface pubescent; mature capitula erect or nodding C. arisanense
– Leaf abaxial surface densely cobwebbed; mature capitula nodding C. hosokawae
9 Leaf abaxial surface densely cobwebbed; mature capitula nodding C. suzukii
– Leaves surface pubescent; mature capitula erect 10
10 Corollas purple; leaves surface shortly hairy C. japonicum var. australe
– Corollas white; leaves surface glabrescent C. japonicum var. takaoense

Species treatments

Cirsium kawakamii Hayata in J. Coll. Sci. Imp. Univ. Tokyo. 159. 1911.

Figs 4B, 5B, 6B, 7B

Type

TAIWAN. Mt. Morrison, ca. 3000 m alt., 20 Oct.1906. T. Kawakami & U. Mori 2279 (holotype: TI!; isotype: TAIF!).

Description

Perennial herbs, stems 0.5−1.8 m tall, without rosette leaves. Leaves pinnatipartite or pinnatisect, 27.5−30.6 cm long and 17.8−20.2 cm wide, U-shaped space between pinnae, smooth, elliptic to broadly elliptic, base truncate to cuneate, apex caudate, pinnae 8.3−10.4 cm long and 17.2−18.6 mm wide, space between pinnae 2.5−3.0 cm, 6−7 pairs. Capitula arranged into racemes or panicles, mature capitula nodding, involucre bowl shaped, 3.4−3.8 cm long and 1.5−2.0 cm wide. Involucre lacking abaxial appendages, inner phyllaries acute apically, outer phyllaries green with indistinct layers, 1.6−1.8 cm long and 1.8−2.4 mm wide, protrusion 6.0−11.0 mm. Florets with white corolla, 2.8−3.1 cm long, corolla lobes 5.2−6.0 mm long and 0.4−0.7 mm wide; 5 synantherous stamens, detached filaments with irregular protuberances, basal caudate extensions, white or brown, anthers 5.4−8.2 mm long, filaments 6.8−8.0 mm long. Stigmas bifid apically, style 2.0−3.4 cm long, ovaries 1.5−2.0 mm long. Achenes oblong, base acute, apex truncate, beige, 4.3−4.9 mm long and 1.7−1.8 mm wide, long tube-shaped beak apically. Pappus 1.3−2.1 cm long forming basal ring, easily shed.

Phenology

Flowering between September and October and fruiting between October and November.

Distribution

Endemic to central-northern Taiwan. Preference for gullies and valleys at 1500−3500 m alt. (Fig. 3).

Chinese name

Yu-shan-ji (玉山薊).

Chromosome number

2n=64 (Fig. 6B).

Palynology

Pollen grains are tricolporate, spheroidal, microreticulate and 31.7−34.5 × 34.3−37.1 μm (P/E ratio: 0.9−1.0). The surface is densely covered with spines that are 2.5−3.2 μm long and 2.1−2.2 μm wide at the base. The distance between spines is 7.6−8.8 μm (Fig. 7B).

Additional specimen examined

TAIWAN. Taoyuan City, Fuxing District, Mt. Lalashan, 1550−1700 m alt., 25 Sept. 1991. C. I Peng 14628 (HAST!). Miaoli County, Tai’an Township, Tunnel of Mt. Shishihshan to Mt. Huoshihshan, 2480 m alt., 18 Sept. 1995. C. M. Wang 1728 (TNM!). Taichung City, Heping District, Mt. Syue to Mt. Chihchiayangdashan, 3300 m alt., 10 Sept. 2014. C. Y. Chang et C. H. Liu 68 (TNM); Mt. Pintian to Mt. Dabajianshan, 2800 m alt., 24°26.52'N, 121°15.83'E, 13 Aug. 2014. C. Y. Chang 264 (TNM); Mt. Syue Trail 8.9 km, 3500 m alt., 24°23.53'N, 121°14.20'E, 3 Oct 2015. C. Y. Chang 774, 1271 (TCF). Nantou County, Ren’ai Township, Nengao cross-ridge historic trail 11 km, 2600 m alt., 24°2.79'N, 121°15.77'E, 29 Jan 2018. C. Y. Chang 1605 (TNM); Chengkung lodge, 3140 m alt., 31 July 2015. C. Y. Chang 654 (TNM); Guandao river, 22 Oct. 1932. S. Sasao s. n. (CHIA!); Mt. Hohwanshan, 3300 m alt., 15 Oct. 1994. Tunghai Collecting Team s. n. (TNM!). Hualien County, Xiulin Township, Sungshiuelou lodge to Dayuling, 3 Aug. 1974. C. N. Lin s. n. (KYO!).

Cirsium tatakaense Y.H.Tseng & C.Y.Chang, sp. nov.

urn:lsid:ipni.org:names:77194985-1
Figs 1, 2, 4A, 5A, 6A, 7A

Diagnosis

Differs from C. kawakamii in having narrower leaf lobes (7.3−11.7 mm), usually more florets, (136)161−308 and phyllaries (111−199), a purplish-red corolla and larger pollen grains (34.2−42.6 × 35.2−44.7 μm).

Type

TAIWAN. Nantou County, Sinyi Township, Highway no. 18, Tataka to Shihshan, 2400 m alt., 23°28.52'N, 120°52.10'E, 3 October 2016. C. Y. Chang 1444 (holotype: TCF; isotype: TNM, PPI).

Description

Perennial herbs, stems 0.5−1.5 m tall, without rosette leaves. Leaves pinnatipartite or pinnatisect, 27.2−34.8 cm long and 16.4−19.4 cm wide, U-shaped space between pinnae, smooth, elliptic to broadly elliptic, base truncate to cuneate, apex caudate, pinnae 6.4−7.5 cm long and 7.3−11.9 mm wide, space between pinnae 2.9−3.0 cm, 4−6 pairs. Capitula arranged into racemes or panicles, mature capitula nodding, involucre bowl-shaped, 3.9−4.0 cm long and 1.7−2.1 cm wide. Involucre lacking abaxial appendages, inner phyllaries acute apically, outer phyllaries reddish-purple with indistinct layers, 1.1−2.2 cm long and 1.8−2.4 mm wide, protrusion 6.3−13.0 mm. Florets with purplish-red corolla, 3.2−3.3 cm long, corolla lobes 4.3−5.3 mm long and 0.4−0.7 mm wide; 5 synantherous stamens, detached filaments with irregular protuberances, basal caudate extensions, light purple or brown, anthers 6.4−6.8 mm long, filaments 7.1−8.1 mm long. Stigmas bifid apically, style 3.0−3.2 cm long, ovaries 1.7−1.9 mm long. Achenes oblong, base acute, apex truncate, beige, 3.0−3.5 mm long and 1.4−1.6 mm wide, long tube-shaped beak apically. Pappus 1.6−1.8 cm long forming basal ring, easily shed.

Phenology

Flowering between August and October and fruiting between September and November.

Distribution

Endemic species of Taiwan. Cirsium tatakaense is located in open areas of cloud forests of vegetation zones from the Quercus to Abies forest zone at alt. 2000−3000 m in central-southern Taiwan (Fig. 3). Based on the geographical climatic regions and vegetation zones (Su 1984, 1985), C. tatakaense is distributed mainly in the central-west inland regions. Cirsium tatakaense has been discovered in sunny environments, such as roadsides and forest margins, concentrated on the upper portions of hills along Provincial Highway no. 18. Miscanthus transmorrisonensis Andersson (Poaceae), Rubus taitoensis Hayata (Rosaceae) and Senecio nemorensis L. var. dentatus (Kitam.) H. Koyama (Compositae) are often discovered with C. tatakaense. Sometimes, C. arisanense Kitam. and C. ferum Kitam. are found near to C. tatakaense; however, no hybrid individual between these species has been observed.

Chinese name

Ta-ta-jia-ji (塔塔加薊).

Etymology

The species epithet tatakaense derives from the type location Tataka in Nantou County.

Notes

This species has in the past been mistakenly identified as C. kawakamii (S. Saito 3477, KYO!; Yamazaki 945, KYO!; C. I Peng 8026, 8936, 11788, 14628, HAST!; K. F. Chung 1053, HAST!; Kawakami & Sasaki s. n., TAIF!; M. L. Weng 1723, TAI!; Y. Kudo & S. Suzuki 300, TAI!; C. C. Hsu 4231, TAI!; C. I Peng 738, TAI!; C. T. Chao et al. 2534, TNM!; C. S. Kuoh 15146, TNM!) or less often as C. arisanense Kitam. (Yamamoto et al. 4142, TAI!; Y. J. Lin 169, PPI!). The earliest record of C. tatakaense was collected by T. Kawakami & S. Sasaki s. n. (TAIF!) at Mt. Morrison (alt. ca. 3000 m) on October 8, 1909. Other specimens collected from 1909−1930 were from Alishan (alt. ca. 2200 m). However, only a few populations remain in Alishan, with the largest population appearing along the Yushan Main Peak Trail from Tataka to Paiyun Lodge (alt. ca. 2800 m). We assume that use in herbal medicine as well as climate change have reduced the population of C. tatakaense.

Chromosome number

2n = 64 (Fig. 6A)

Palynology

Pollen grains are tricolporate, spheroidal, microreticulate and 34.2−42.6 × 35.2−44.7 μm (P/E ratio: 0.9−1.0). The surface of the pollen is densely covered with spines that are 3.2−5.1 μm long and 4.2−5.6 μm wide at the base. The distance between spines is 7.5−10.6 μm (Fig. 7A).

Conservation status

Cirsium tatakaense is distributed in central-southern Taiwan, with a population of more than 1000 mature individuals. Its habitats are mainly located in high and sunny mountain areas and many of them are difficult to locate. Therefore, following the International Union for Conservation of Nature (IUCN) Categories and Criteria (IUCN 2014), we regard this species as Least Concern. However, long-term monitoring of its population is still required.

Additional specimens examined (paratype)

TAIWAN. Nantou County, Sinyi Township, Highway no. 18, Tataka to Shihshan, 2400 m alt., 23°28.52'N, 120°52.10'E, 3 Oct. 2016. C. Y. Chang 1442 (TCF); same loc., 3 Oct. 2016. C. Y. Chang 1443 (TCF); same loc., 12 Sept. 2012. C. T. Chao et al. 2534 (TNM!); Tunpu Hot Spring to Kuankao, 1300−2600 m alt., 3 July 1985. C. I Peng 8026 (HAST!). Chiayi County, Alishan Township, Alishan, 25 Dec. 1928. Y. Kudo & S. Suzuki 300 (TAI!); Tatachia saddle to Paiyunshanchuang, 2700−3000 m alt., 9 Nov. 1985. C. I Peng 8931 (HAST!); Tatachia saddle to Mt. Yushanchienfu, 2700−3100 m alt., 11 Nov. 1990. P. J. Wu et al. s. n. (TNM!). Kaohsiung City, Tauyuan District, Kuaiku to Yakou, 2600 m alt., 23°16.03'N, 120°56.24'E, 21 Oct. 2014. C. Y. Chang 160 (TNM); Gingzin bridge, 14 Sept. 1999. C. S. Kuoh 15146 (TAN!). Taitung County, Yanping Township, Yenping forest-road, 1500−1800 m alt., 2 July 2006. Y. J. Lin 169 (PPI!).

Figure 1. 

Line drawings of Cirsium tatakaense Y.H.Tseng & C.Y.Chang A habit B root C leaf D capitula E inner phyllary E’ middle phyllary E” outer phyllary F floret F’ floret (pappus removed) G synantherous H style branches I achene. Voucher: C. Y. Chang 1442 (TCF).

Figure 2. 

Cirsium tatakaense Y.H.Tseng & C.Y.Chang A habitat B habit C seedling D variations of leaves E inflorescences F capitula G floret; (right-side pappus removed) H achene with pappus I achene.

Figure 3. 

Distribution map of Cirsium tatakaense Y.H.Tseng & C.Y.Chang (star); C. kawakamii Hayata (triangle); and C. arisanense Kitam. (circle) of Taiwan.

Figure 4. 

Holotypes of the three species of Cirsium sect. Onotrophe in Taiwan. A C. tatakaense Y.H.Tseng & C.Y.Chang, C. Y. Chang 1444 (TCF) B C. kawakamii Hayata, T. Kawakami & U. Mori 2279 (TI!) C C. arisanense Kitam., S. Kitamura s.n. (KYO!).

Figure 5. 

Comparison of the morphological characters amongst the species of Cirsium sect. Onotrophe in Taiwan. A C. tatakaense Y.H.Tseng & C.Y.Chang B C. kawakamii Hayata C C. arisanense Kitam.: 1 leaf 2 capitula 3 inner phyllary 3’ middle phyllary 3” outer phyllary 4 floret 4’ floret (pappus removed) 5 synantherous 6 style branches 7 achene 7’ achene with pappus.

Figure 6. 

Chromosome number of the two species of Cirsium subsect. Nipponocirsium Kitam. in Taiwan. A 2n = 64, C. tatakaense Y.H.Tseng & C.Y.Chang B 2n = 64, C. kawakamii Hayata. Scale bar: 5 μm.

Figure 7. 

Comparison of the pollen morphology of the three species of Cirsium sect. Onotrophe in Taiwan. A C. tatakaense Y.H.Tseng & C.Y.Chang B C. kawakamii Hayata C C. arisanense Kitam.: 1 polar view 2 equatorial view 3 colporate view. Scale bar: 30 μm.

Acknowledgements

We thank Dr. Ching-I Peng (彭鏡毅) and Dr. Chien-Ti Chao (趙建棣) for providing useful suggestions and we also thank Mrs. Chia-Hao Liu (留嘉豪), Sheng-Hua Tseng (曾勝華), Tai-Yuan Chang (張台員) and Ms. Pi-Ling Wang (王碧玲) for field assistance. This manuscript was edited by Wallace Academic Editing.

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